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THE YEAR IN AND , 2009

Climate Change Adaptation Strategies for Management and Conservation Planning Joshua J. Lawler College of , University of Washington, Seattle, Washington

Recent rapid changes in the Earth’s climate have altered ecological systems around the globe. Global warming has been linked to changes in physiology, phenology, species distributions, interspecific interactions, and disturbance regimes. Projected future cli- mate change will undoubtedly result in even more dramatic shifts in the states of many ecosystems. These shifts will provide one of the largest challenges to managers and conservation planners. Managing natural resources and ecosystems in the face of uncertain climate requires new approaches. Here, the many adaptation strategies that have been proposed for managing natural systems in a changing climate are reviewed. Most of the recommended approaches are general principles and many are tools that managers are already using. What is new is a turning toward a more agile management perspective. To address , managers will need to act over different spatial and temporal scales. The focus of restoration will need to shift from historic species assemblages to potential future ecosystem services. Active adaptive management based on potential future climate impact scenarios will need to be a part of everyday operations. And triage will likely become a critical option. Although many concepts and tools for addressing climate change have been proposed, key pieces of information are still missing. To successfully manage for climate change, a better un- derstanding will be needed of which species and systems will likely be most affected by climate change, how to preserve and enhance the evolutionary capacity of species, how to implement effective adaptive management in new systems, and perhaps most importantly, in which situations and systems will the general adaptation strategies that have been proposed work and how can they be effectively applied.

Key words: adaptation; adaptive management; climate change; conservation planning; management; scenario planning; triage

◦ Introduction jected to rise between 3 and 12 Cbytheendof the century (IPCC 2007b). Precipitation pat- Over the past century, global average annual terns are also projected to change, although temperatures have risen 0.7◦C (IPCC 2007b). the direction, magnitude, and confidence sur- In the Arctic, temperatures have risen at ap- rounding precipitation projections vary by re- proximately twice that rate. This trend is very gion and season. likely to continue into the future, as global aver- These changes have profound implications age surface temperatures are projected to rise for the Earth’s natural systems. Recent climatic between 1.1 and 6.4◦C by 2100, and temper- changes have been linked to decreases in snow- atures at the high northern latitudes are pro- pack (Groisman et al. 2001; Mote 2003), in- creases in the frequency and severity of large wildfires (Westerling et al. 2006), and rising sea levels (IPCC 2007b). These changes in Address for correspondence: Joshua J. Lawler, College of Forest Re- sources, University of Washington, Box 352100, Seattle, WA 98105. turn have the potential to alter the timing and [email protected] magnitude of stream flows, the structure and

The Year in Ecology and Conservation Biology, 2009: Ann. N.Y. Acad. Sci. 1162: 79–98 (2009). doi: 10.1111/j.1749-6632.2009.04147.x C 2009 New York Academy of . 79 80 Annals of the New York Academy of Sciences composition of vegetation communities, and sive species will emerge, and ecosystem func- the nature of coastal systems. As temperatures tions will be altered. and sea levels continue to rise, many ecosys- , corals, and other organisms have tems will undergo significant changes. Coastal clear physiological responses to climate change. wetlands will be inundated, alpine zones will Increased atmospheric CO2 concentrations shrink, and some wetlands, ponds, and lakes can result in increased -use efficiency in will dry up (IPCC 2007a). plants. Because different species will respond Many ecological systems are already show- differently to increased CO2, differential in- ing the effects of recent climatic changes creased water-use efficiencies will likely result in (Walther et al. 2002; Parmesan and Yohe 2003; shifts in competitive relationships and changes Root et al. 2003; Parmesan 2006). The most in communities (Policy et al. 1993). Many well documented changes include changes in corals are extremely sensitive to changes in tem- phenology, species distributions, and physiol- perature. Increases of just a few degrees for even ogy. Recent phenological changes have been a short period can result in bleaching (a loss of observed in many different ecological systems the coral’s symbiotic zooxanthellae and their (Sparks and Carey 1995). Spring events, for ex- photosynthetic pigments). Extensive bleaching ample, have been occurring 2.3 days earlier events have occurred during the past 20 years per decade over the last century (Parmesan in several regions around the globe (West and and Yohe 2003). Plants are flowering and fruit- Salm 2003). Several species of fish also have ing earlier (Cayan 2001), birds are laying eggs well-documented thermal thresholds for sur- earlier (Brown et al. 1999; Crick and Sparks vival at different stages (e.g., McCullough 1999), and some amphibians are mating ear- 1999; Moyle 2002). Even small changes in tem- lier (Beebee 1995; Gibbs and Breisch 2001). perature have the potential to affect population Changes in phenology have the potential to dynamics and habitat use for many species. decouple interdependent ecological events, re- Managing natural systems in the face of such sulting in changes in species interactions, com- widespread change is a daunting task. Perhaps munity composition, and ecosystem function- the largest challenge for mangers is making de- ing (Stenseth and Mysterud 2002). cisions based on limited and often highly un- The paleoecological record indicates that certain projections of future climate impacts species have shifted their geographic ranges (Lawler et al. in press-b). Over the past 10 to in the past in response to changes in climate 20 years, researchers have begun to suggest (Brubaker 1989; Davis and Shaw 2001). More ways that managers and planners can begin recent records indicate that species have also to address climate change. Here, I summa- shifted their distributions in response to re- rize these recommendations. I begin with an cent climate change. Many species of plants, overview of the general, largely conceptual, rec- birds, butterflies, and amphibians have shifted ommended adaptation strategies. I go on to their distributions in patterns and at rates that describe some of the more specific suggestions are consistent with recent climatic changes that have been made for addressing climate (Parmesan 2006). In general, these species are change in freshwater, marine, and terrestrial shifting their ranges upward in elevation and systems. Although many of the recommended poleward in latitude (Parmesan et al. 1999; approaches for addressing climate change are Thomas and Lennon 1999; Seimon et al. 2007; already used to manage resources and protect Lenoir et al. 2008). In a few cases, popula- , effectively implementing these ap- tion and even species extinctions have been at- proaches will require new perspectives. I con- tributed to recent climatic changes (Pounds et al. clude with a brief discussion of the most press- 1999). As species move in response to climate ing research needs for successfully developing change, new communities will form, new inva- and implementing adaptation strategies. Lawler: Climate Change Adaptation Strategies 81

General Strategies for Addressing toxicity of pesticides or the infection rates and Climate Change severity of diseases (Kumaraguru and Beamish 1981). Likewise climate change may increase The vast majority of the proposed strategies competitive pressure from invasive species, as for managing resources in a changing climate some invasive species may benefit from in- are general concepts. These concepts can be creased atmospheric CO2 concentrations or loosely grouped into three basic types of strate- changes in temperature or precipitation, allow- gies: those promoting resistance, resilience, and ing them to spread and/or outcompete native change (Millar et al. 2007). Resistance is the abil- species (Dukes and Mooney 1999; Schlesinger ity of a system to remain unchanged in the face et al. 2001; Zavaleta and Royval 2001; Rahel of external forces. Resilience can be defined as and Olden 2008). the ability of a system to recover from pertur- Environmental stresses may also reduce bations (Holling 1973). A resilient system will the resilience of individuals and populations change in response to external forces but will to climate change. For example, Drosophila return to its original state. With respect to cli- melanogaster exposed to parasitic attacks while mate change, systems that are more resilient are in the larval stage were more susceptible to those that are better able to adapt to changes desiccation that those not exposed to para- in climate. Resilient systems will continue to sitic attacks (Hoang 2001). Similarly,streamside function, albeit potentially differently, in an al- salamanders, Ambystoma barbouri, exposed to the tered climate. Less resilient systems will likely herbicide atrazine were more susceptible to undergo messy transitions to new states, result- desiccation than were unexposed salamanders ing in the loss of ecosystem functioning, popu- (Rohr and Palmer 2005). Removing or reduc- lations, or even species. Strategies that promote ing existing environmental stresses and threats change are those designed to help move a sys- to populations or species will, in some cases, tem from one state to another. The most com- enhance their resilience to climate change. monly recommended strategies for promoting resistance, resilience, and change are briefly Expanding Reserve Networks discussed below. Protected areas are arguably one of the best Removing Other Threats and Reducing ways to conserve biodiversity.However, climate Additional Stresses change will challenge the ability of the current reserve network to provide protection when the Perhaps the most obvious approaches to in- climate shifts so much that plants and animals creasing resilience are the removal of other, no longer thrive where their current reserves non-climate-related threats to a species or sys- are located (Peters and Darling 1985). Many tem and reducing other stresses on species. De- researchers have suggested expanding reserve creasing the impact of exotic species, habitat networks to give systems and species room to loss and fragmentation, overharvest, and other move and places to go (Halpin 1997; Shafer threats generally results in larger populations 1999). Increasing redundancy in the reserve that will likely be better able to absorb per- network can also increase resilience by provid- turbations (Rogers and McCarty 2000; Noss ing more opportunities in different places or 2001; Soto 2001; Hansen et al. 2003). Not only chances in which species or communities might do other threats reduce the ability of a popula- persist. Some have recommended increasing tion or system to respond to or to absorb new the size of existing reserves, adding buffers impacts, but in many cases, climate change may around existing reserves, and adding larger re- exacerbate the effects of other threats. For ex- serves to reserve networks (Halpin 1997; Shafer ample, increases in temperature may increase 1999; Noss 2001) (Fig. 1A). Larger reserves are 82 Annals of the New York Academy of Sciences

Figure 1. Proposed strategies for augmenting existing reserve networks to address climate change. Additional reserves can be placed (A) to enlarge existing reserves, (B) to span climatic or edaphic gradients, (C) to facilitate directional species movements in response to increasing temperatures, and (D) to help connect existing reserves. Existing reserves are represented by darker shapes, and new reserves are represented by lighter shapes. likely to preserve a greater diversity of envi- and placing new reserves between existing re- ronmental conditions and allow for movement serves to facilitate connectivity (Fig. 1D). within the reserve. Coastal systems in partic- Ideally, ecological forecasting could be used ular will require large reserves that extend to identify sites that would best protect biodi- inland, allowing species to shift as sea levels versity in a changing climate (Hannah 2008). rise. Dynamic global vegetation models (DGVMs; Simply developing more and larger reserves e.g., Cramer et al. 2001; Bachelet et al. 2003) and might not be enough if they are not located in climate-envelope models (Pearson and Dawson the right places. One more strategic possibility 2003) have been used to assess the ability of entails locating reserves so that they capture the reserves or reserve networks to protect species most potential for habitat heterogeneity under under different climate-change scenarios (Scott any climate scenario. Such areas would cover et al. 2002; Burns et al. 2003; Araujo´ et al. 2004). diverse topographic, edaphic, and hydrologic A few studies have used such models to sug- conditions (Halpin 1997). Placing large, long gest reserve networks that would be resilient reserves across biotic transition zones such as to climate change. For example, Hannah et al. ecotones may allow for the continued protec- (2007) used climate-envelope models to project tion of that transition as it shifts with climate species’ range shifts in three different regions change (Fig. 1B). Others have suggested plac- of the globe in response to climate change and ing reserves at the poleward edge of species then selected reserves that adequately protected ranges (Shafer 1999), arranging reserves longi- those species in the future. Likewise, Williams tudinally (Pearson and Dawson 2005) (Fig. 1C), et al. (2005) used climate-envelope models to Lawler: Climate Change Adaptation Strategies 83 identify potential corridors that would allow for a given species that protect a wide range of movement between current ranges and species’ climatic conditions within that species’ range potential future ranges. Such approaches rely (Pyke et al. 2005; Pyke and Fischer 2005). heavily on the ability of the models to pre- dict species’ responses to climate change. Given Increasing Connectivity both the in projected future cli- mates and the uncertainty inherent in most In the past, species have moved across con- relevant ecological forecasting approaches (e.g., tinents as climates changed and ad- Thuiller 2004; Lawler et al. 2006), use of these vanced and retreated (Davis and Shaw 2001). models for the selection of reserves will require, One of the biggest differences between those at the very least, that meaningful uncertainty historic periods and today is that humans estimates are evaluated in conjunction with, or have dramatically altered the Earth’s surface. incorporated into, the future projections. A sec- Agricultural , roads, dams and water di- ond limitation of this approach is that data will versions, urban areas, and residential develop- be lacking for the vast majority of biodiversity, ment all act as barriers to movement for some and reserve selection based on this approach is species. These barriers will make it difficult for likely to be biased toward well-studied species many species to move from areas that become with ample data. unsuitable or to occupy new climatic zones or One alternative to using predicted shifts in habitats that emerge in the future. Using a com- species’ ranges or changes in vegetation is to bination of statistical climate-envelope model- use projected changes in climate alone or in ing and mechanistic dispersal models, Iverson conjunction with static elements of the environ- et al. (2004) explored the potential of five eastern ment to determine where conditions are likely American tree species to colonize newly emerg- to be more or less constant. Saxon et al. (2005) ing climatic space over a 100-year period. None used a combination of edaphic conditions and of the five species was able to colonize more projected future climates to identify environ- than 15% of the areas projected to be suitable ments that would likely change more and those under future climates. Such work implies that that would likely change less. Reserves could the number of species that are able to move suc- be located in areas projected to experience less cessfully in response to today’s anthropogenic change, or in areas that connected shifting en- climatic changes will be a small fraction of the vironments. Such an approach relies on pro- number that were able to move in response to jected future climatic conditions, and thus is historic climate shifts. still imbued with uncertainty, albeit less uncer- The highly fragmented nature of today’s tainty than approaches that involve additional has led many conservation biolo- climate-envelope or DGVM modeling. gists to promote increasing connectivity among Another alternative that has been proposed protected areas to enhance movement in a is to base reserve selection on the underlying changing climate (Shafer 1999; Noss 2001; edaphic conditions and/or current climatic or Hulme 2005; Welch 2005). Most of the research bioclimatic gradients. Bedrock, , topogra- and discussion on connectivity has focused on phy, and climatic gradients largely define the corridors, although some work has ad- natural distribution of flora, fauna, and ecosys- dressed corridors for plants (e.g., Williams et al. tems. By protecting diverse combinations of 2005). Almost all of this work has taken a these factors, it may be possible to preserve the species-by-species approach to corridors. Such ecological stage on which new players (species) an approach is justified, given that corridors, will find themselves in future climates. One like habitat, are species-specific concepts. The such approach that captures current climatic ability to move through a corridor depends on gradients would be to select multiple reserves species-specific behavior and habitat affinities. 84 Annals of the New York Academy of Sciences

Given that many species, with diverse habitat return a system to historic or predisturbance requirements and dispersal abilities, will need conditions (Swetnam et al. 1999). In some cases, to move in response to climate change, species- this may still be a viable option. However, based corridor approaches may not be ade- for many systems, climate change will make it quate or feasible (Hulme 2005). costly if not impossible to recreate past ecolog- Two additional approaches to increasing ical states. Changes in atmospheric CO2 con- connectivity have been proposed. First, as men- centrations will likely alter competitive relation- tioned earlier in this chapter, small stepping- ships between plant species in multiple ways stone reserves can be placed between larger (Schlesinger et al. 2001). For example, increased preserves to facilitate movement (Shafer 1999) water-use efficiency due to increased atmo- (Fig. 1D). This approach may provide connec- spheric CO2 concentrations may allow trees tivity for a more general group of species than to move into more arid environments, shift- species-specific corridors, but small reserves will ing shrublands to woodlands and woodlands be limited to a small range of environmental to closed-canopy . Trying to maintain conditions, and thus may only provide habi- a shrubland in such an area of transition will tat for select and potentially small groups of be costly and potentially ineffective. Second, species. Similarly, new reserves, whether they much current restoration takes a species-based are small or large, can be placed in close prox- approach, focusing on restoring species com- imity to existing reserves to facilitate move- position. Successfully restoring specific species ment (Halpin 1997). The second approach is to assemblages will require relatively accurate pre- manage the lands or between protected dictions of which sites will be suitable for which areas in ways that allow the most species to species in the future and how those species move through these spaces. Such approaches will interact under projected climatic condi- have been referred to as softening or man- tions. Given the uncertainty inherent in fu- aging the matrix (Franklin et al. 1992; Noss ture climate projections, our limited knowledge 2001). Some combination of matrix manage- of species-specific responses to climate change, ment, stepping-stone reserves, and corridors and the inherent in most eco- will likely allow the most movement in response logical forecasting tools, it is unlikely that we to climate change. will soon have future predictions that are ac- curate enough to successfully describe specific Restoring Habitat and System Dynamics species assemblages at a given site in the distant future. Clearly, the restoration of ecosystem func- Harris et al. (2006) suggest a shift away tioning and habitat in degraded areas plays a from restoring historic conditions and specific key role in increasing resilience for systems and species assemblages. As an alternative to these species. Many have highlighted the need to re- traditional approaches, they propose restor- store functioning ecosystems to address climate ing process instead of structure. By focusing change (Hartig et al. 1997; Mulholland et al. on ecosystem services instead of species com- 1997; Joyce et al. 2008; Peterson et al. 2008). position, ecosystems can be managed in a However, climate-induced changes in hydrol- more flexible way in which species assemblages ogy, disturbance regimes, and species distribu- change with changing climates, but ecosys- tions will make restoration goals moving tar- tem functioning—although the nature of those gets, thereby challenging the way restoration is functions might change—is preserved. Sev- typically done (Harris et al. 2006). eral suggestions have been made for restor- Climate change calls into question two of ing disturbance regimes, river and stream- the basic tenants of restoration (Harris et al. flow regimes, and wetland hydrology (Hartig 2006). First, most current restoration aims to et al. 1997; Noss 2001; Harris et al. 2006). Lawler: Climate Change Adaptation Strategies 85

Concentrating on the abiotic aspects of systems of highly uncertain systems. Thus, it is poten- will help to prepare sites and systems for new tially an ideal approach for dealing with the un- conditions and new sets of species. certainties surrounding future climatic changes Although ecological models may not yet be and future climate impacts (Arvai et al. 2006). able to provide accurate enough predictions of Adaptive management can be passive or active future species assemblages to allow managers (Walters 1986; Walters and Holling 1990). Pas- to concentrate all of their efforts on a few key sive adaptive management generally involves species, models are currently accurate enough building a management strategy based on his- to inform restoration efforts. Climate-envelope toric data and then altering that strategy with models and DGVMs can provide an estimate of new data as the system is monitored over how the species composition at a site might shift time. Active adaptive management involves over time, giving managers an idea of the types conscious experimentation, generally exploring of new species that they might incorporate into the outcomes of multiple management strate- restoration actions and the species for which gies. Both types of adaptive management will continued preservation at a site might be futile. likely be needed to address climate change. Simulation models can also be used to explore Applying adaptive management to address the potential effects of restoration efforts in a climate change will be an iterative multistep changing climate. Battin et al. (2007) integrated process (Kareiva et al. 2008). First, such a pro- the results of climate models, -use models, cess will involve assessing the potential impacts hydrology models, and populations models to of climate change on a system. This would investigate the potential effects of restoration necessarily be a comprehensive assessment in efforts for Chinook salmon under two different which as many of the potential ramifications of climate-change scenarios. Higher stream tem- changes in climate, including changes to distur- peratures, higher peak winter flows, and lower bance regimes, hydrology, species composition, flows during spawning are likely to lead to de- resources, phenology, and interspecific in- creases in salmon populations. Because changes teractions are all considered. Second, manage- in flow regimes had the largest effect at higher ment actions will need to be designed to ad- elevations and restoration efforts were concen- dress these impacts. These actions should be trated at lower elevations, their model indicated seen as experiments. In some cases, in which that the combined effect of climate change the scale and the nature of the problem are and restoration would shift salmon breeding amenable, it will be possible to conduct multi- to lower elevations. ple experiments to explore competing hypothe- sized effects of climate change or to account for Adaptive Management different potential shifts in climate. However, for large-scale management problems, a pas- Adaptive management is often cited as a sive adaptive management approach will be critical approach to addressing climate change necessary. Third, the system will need to be (Millar et al. 2007; Kareiva et al. 2008; Lawler monitored for both climatic changes and po- et al. in press-b). Adaptive management in- tential system responses. Finally, management volves an iterative process in which managers strategies will need to be reevaluated and po- learn from experimental management actions tentially redesigned before the cycle is repeated (Holling 1978; Walters and Hilborn 1978). (Kareiva et al. 2008). Management actions are applied as experi- Although adaptive management is a very ap- ments, the system is monitored, and actions are pealing concept and it is often written into then potentially changed to address changes management plans, it has been implemented in the state of the system. In theory, adap- in relatively few instances (Walters 1997). tive management allows for the management Two of the better known examples include 86 Annals of the New York Academy of Sciences water management in the Florida Everglades with this information, predicting invasion is dif- (Walters et al. 1992) and the management of ficult. Second, determining where to transplant flows for the Glenn Canyon Dam (National species requires an understanding of how en- Research Council 1999). Some of the barriers vironments will change in the future and what to successfully implementing adaptive manage- new areas will provide suitable habitat. As dis- ment include the lack of institutional flexibility cussed earlier in the chapter, projections of fu- and capacity, the perceived of failure, high ture climatic conditions and ecological forecasts degrees of uncertainty, and large spatial and based on those projections are still relatively long temporal scales of management (Gregory uncertain. To be useful for assessing potential et al. 2006). All of these factors will challenge sites for translocations, these projections and adaptive management as a tool to address cli- forecasts will need to incorporate these uncer- mate change. Nonetheless, it is still likely to be tainties. Given these potential barriers to imple- one of the best tools managers and scientists menting translocations, a framework for assess- have to address climate change and to learn ing the feasibility of, and risks associated with, about its effects. a given translocation is essential. Hoegh Guldberg et al. (2008) recently Translocations proposed just such a framework (Fig. 2). Their proposed framework involves several basic Even with increased connectivity between questions and management options. The first protected areas, some species will need to be question asks what the of extinction or pop- moved to prevent extinction. Species with lim- ulation decline is likely to be under future pro- ited dispersal abilities and small, isolated ranges jected climates. If the risk is low, assisted mi- may have trouble tracking shifting habitats or gration is not likely to be needed. If the risk may be left without suitable habitat altogether. is moderate, the authors suggest enhancing re- For at least some of these species, translocations silience by increasing connectivity, may be the only solution (Bartlein et al. 1997; reducing non-climate-related threats, and in- Honnay et al. 2002). Although recommenda- creasing genetic diversity. If the risk is high, tions for translocation are not new (Peters one asks whether translocation and establish- and Darling 1985; Orians 1993), conservation ment is technically possible (the second ques- biologists have generally been reluctant to tion in the framework). If not, actions may still tackle the issue of the purposeful movement be taken to help create suitable habitat in new of species outside of their known native range. areas in hopes that organisms will get there Such translocations raise critical ecological and on there own. If translocation is possible, one ethical questions that will need to be addressed asks the third question—whether the benefits before attempts to relocate species are made of a translocation outweigh the potential eco- (Hunter 2007; McLachlan et al. 2007). logical and socioeconomic costs. If the bene- Given the impacts that invasive species have fits outweigh the costs, translocations can be on ecological and economic systems, translo- undertaken. cations may have costly ramifications even if well researched, planned, and executed. McLachlan et al. (2007) discuss two of the basic Specific Recommendations barriers to implementing translocations. First, for Addressing Climate Change it is difficult to predict which potential candi- dates for translocation will potentially become The strategies discussed in the preceding sec- invasive. Although research on invasive species tions are all basic concepts or general recom- has produced some information about com- mendations. Here, I provide a few examples of mon traits of invasive species, in general, even more specific actions that have been proposed Lawler: Climate Change Adaptation Strategies 87

Figure 2. Framework for planning potential species translocations in a changing climate. [This figure was adapted with permission from Hoegh-Guldberg et al. (2008) and AAAS.] for addressing climate change in freshwater, changes in temperature will result in reduced marine, and terrestrial systems. dissolved oxygen concentrations and changes in species composition. Freshwater Systems Rising water temperatures will result in shifts in species distributions, including the potential In response to climate change, freshwater loss of cold-water fish from some lower stream systems are expected to experience changes reaches and from other streams all together in temperature, flow, evaporation rates, water and expansions of the ranges of warm-, and species composition (Frederick and fish (Carpenter et al. 1992; Eaton and Scheller Gleick 1999; Poff et al. 2002). In many mon- 1996). Riparian restoration has been proposed tane systems, reduced snowpack and earlier as one method to reduce stream temperatures spring melts will result in changes in the tim- and create cool water refugia (Palmer et al. ing and intensity of spring and summer stream 2008; Scott et al. 2008). Riparian restoration flows (Barnett et al. 2005; Milly et al. 2005). projects would also help to provide connec- Wetlands, particularly those that are depen- tivity among some terrestrial systems. Protect- dant on precipitation, are likely to be some ing headwaters and identifying and protecting of the most susceptible to climate change of existing thermal refugia will also enhance the all aquatic systems (Burkett and Keusler 2000; ability of cold-water fish to persist as tempera- Winter 2000). Increased evaporation rates and tures rise (Hansen et al. 2003). 88 Annals of the New York Academy of Sciences

A number of management actions have been of elevated water temperatures. Another ap- suggested for dealing with increases and de- proach to protecting coral reefs involves iden- creases in flows (Palmer 2008). Some of these tifying resistant and resilient coral populations include channel reconfiguration, dam removal that are less susceptible to higher temperatures or retrofit, floodplain restoration, dam-based (Hansen et al. 2003; West and Salm 2003). flow management, and bank stabilization. For These populations may be able to recolonize example, creating wetlands and off-channel bleached reefs, or may be transplanted in at- basins for water storage during times of extreme tempts to restore damaged reefs. flows may prevent excess water from reach- As in other systems, reducing other threats ing reservoirs and reduce downstream flows to species or ecosystems will likely enhance the (Palmer et al. 2008). The removal of sediment resilience of marine systems. Reducing fishing from reservoirs may also increase water storage pressure and damaging fishing techniques such capacity in the short term (Palmer et al. 2008). as dynamite and cyanide fishing will increase Water releases from dams and transporting fish the resilience of fish stocks and reefs (Hansen may be necessary short-term solutions in times et al. 2003). Reducing nutrient pollution will of drought or extreme low flows (Palmer 2008). likely reduce the frequency and extent of harm- Finally, reducing water extraction will be a key, ful algal blooms that are a combined result of although controversial, approach to maintain- increased nutrients and increased ocean tem- ing flows (Hansen et al. 2003). peratures (Mudie et al. 2002). Decreasing load- ing of nutrients and other pollutants will require Marine Systems changing land-use practices in coastal water- sheds and in many cases over much larger areas Although marine systems are less studied (Hansen et al. 2003). than terrestrial systems and the Intergovern- mental Panel on Climate Change (IPCC) Terrestrial Systems has documented fewer significant biological changes in marine systems, many of those A common recommendation for enhancing changes can be linked to climate change the adaptive capacity of terrestrial systems, par- (Richardson and Poloczanska 2008). As in ticularly forested systems, is to broaden the freshwater and terrestrial systems, species genetic variability and the species diversity of movements, phenological shifts, and physiolog- managed sites (Harris et al. 2006; Millar et al. ical effects of climate change will have signifi- 2007). Instead of using seed or individuals from cant effects in marine systems (Perry et al. 2005; local or even regional populations, it may be Portner and Knust 2007). Marine species and advantageous to maximize the genetic diver- systems, however, face an additional challenge sity at a restoration or reintroduction site by as increased CO2 concentrations continue to taking genetic material from a broader range acidify the oceans (Ruttimann 2006; Guinotte of locations within a species, range. and Fabry 2008). Many researchers have called for aggressive Increasing ocean temperatures have resulted forest-management practices to address climate in the bleaching of corals around the world. A change in forested systems. For example, widely number of specific adaptation strategies have spaced thinning and shelterwood cuts may al- been proposed to address the loss of corals low forest stands to withstand increased insect (Hansen et al. 2003). One such strategy involves outbreaks and fires (Dale et al. 2001; Joyce et al. the reduction of synergistic stressors such as 2008). Prescribed burning can be used to re- high irradiance. Shading and water disruption duce fuel loads, and hence the risk of catas- (with sprinklers) are two methods that are being trophic fire (Spittlehouse and Stewart 2003; explored to reduce irradiance during periods Scott et al. 2008). Furthermore, aggressive site Lawler: Climate Change Adaptation Strategies 89 preparation has been proposed to enhance re- and managing for dynamic processes and com- generation after disturbances (Spittlehouse and munities. Other shifts in perspectives that will Stewart 2003). be required to address climate change include Manipulative management strategies have broadening the scale of management, engaging been suggested for other systems as well. For ex- in scenario-based planning and management, ample, moderate grazing has been proposed to and embracing triage as a necessary tool. increase the hydroperiod in vernal pools threat- ened by increasing temperatures and decreas- Scale ing precipitation (Pyke and Marty 2005). The placement of snow fences has been proposed Effectively managing resources in a changing to increase snow pack in areas where sensi- climate will require taking a broader spatial and tive alpine plant communities are threatened temporal perspective (Franklin et al. 1992; Scott by reduced snowpack (Hansen et al. 2003). For et al. 2002; Welch 2005; U.S. Envioronmental a wide variety of systems, invasions by nonna- Protection Agency 2008). Instead of consider- tive species may be minimized through vigi- ing a population, community, or ecosystem in lance, early detection, and aggressive removal isolation, it will be essential to manage these (Hansen et al. 2003; Baron et al. 2008). within a regional or even continental context. For example, it will be necessary to consider the relative location of a population within the New Perspectives geographic range of the species and to coordi- nate the management of that population with None of the general or specific strate- others throughout the range. Given changes gies mentioned here is new. For example, in climatic conditions, it may be necessary to even translocations are not without historic abandon efforts to manage a population at one analogs. Reintroductions are merely transloca- edge of the species’ range and to shift efforts to tions within the historic range of a species. Fur- other populations elsewhere in the range. Be- thermore, the introduction of species for biolog- cause management strategies and practices will ical control should, theoretically,evoke a similar need to cross both land-ownership and political set of precautions that need to be taken when borders, they will require both intergovernmen- species are moved to address climate change. tal and interagency coordination (Soto 2001; In fact, most of the general adaptation strate- Hannah et al. 2002). In particular, some re- gies and many of the specific actions described searchers have called for new administrative earlier are the basic accepted strategies for pro- structures such as interagency teams or pro- tecting biodiversity in general. How, then, will grams with a mandate to address climate management need to change in the face of cli- change (Kareiva et al. 2008). mate change? Although many of the traditional strategies Scenario-Based Planning for protecting biodiversity will be critical for ad- dressing climate change, applying these strate- The uncertainty inherent in future climate- gies will require a new perspective. Thus, most change projections means that it will often be of the necessary change in management will risky to manage for one anticipated future cli- revolve around how established strategies are mate. Current practice often involves manag- applied. For example, as discussed earlier in ing for a set goal, for example, a population size, the chapter, restoration will be a critical tool a particular forest structure, an allowable sed- for addressing climate change. However, suc- iment load, or a specific species composition. cessful restoration in a changing climate will One or more strategies are then developed to require largely abandoning historic conditions attain that set goal. In certain climates, a given 90 Annals of the New York Academy of Sciences goal may be unattainable. Climate change will Regardless of which climate scenario was ap- force managers and planners to evaluate mul- plied by Battin et al. (2007), restoration and tiple potential scenarios of change for a given protection of lowland salmon habitats yielded system and then to develop alternative manage- significant benefits. Attempting to identify these ment goals and strategies for those scenarios. robust management options may prove more Scenario-based planning has its origins in fruitful than trying to determine an optimal military theory, and it has been used to explore management strategy. planning strategies in a variety of disciplines Scenario planning can be incorporated when future conditions are uncertain (Peterson into adaptive management to help managers et al. 2003). The IPCC uses a scenario-based address climate change. Whereas a more tra- approach to exploring the range in potential ditional approach to active adaptive man- future climates given different greenhouse-gas agement might involve simultaneously testing emissions scenarios (Nakicenovic et al. 2000). several alternative management strategies to The different scenarios make different assump- attain a set goal, active adaptive management tions about human population growth, eco- for climate change will require testing several nomic and social cooperation, advances and different management strategies designed to acceptance of new technologies, and consump- attain different goals under different climate- tion. Applying these scenarios results in dif- change scenarios. Developing the different sce- ferent projected concentrations of greenhouse narios and goals will require knowledge of the gasses, which, in turn, result in different degrees range of plausible future climate-change pro- of warming and different changes in precipita- jections and some estimate of how those dif- tion (IPCC 2007b). Alternative future scenarios ferent climatic changes will affect the system have also been used to evaluate the impacts of or species in question. This type of scenario- alternative development strategies on biodiver- based approach will require managers to think sity (White et al. 1997; Schumaker et al. 2004). in parallel, in essence, planning for a system In these studies, land-use scenarios were de- with several potential future states. veloped based on preferences for conservation and urban and suburban development. Mod- Triage els were then used to evaluate the potential im- pact of the different scenarios on population Given the relative scarcity of funding for persistence. conservation and management of natural Battin et al. (2007) explored the effects of two resources, prioritizing management needs is different climate-change scenarios and three already an integral part of the organization- different restoration scenarios on salmon pop- wide planning of most government agencies ulations. The two different climate scenarios and environmental nongovernmental organi- were projected to increase peak flows during zations. Climate change will likely stretch those the salmon incubation period by 7% and 28%, funding resources even thinner. With many sys- respectively. The six different scenario combi- tems, species, and sites requiring active adap- nations resulted in different changes in popula- tive management to address climate change, tion size, ranging from a 40% decrease to a 19% managers and planners will have to make dif- increase, depending on the scenario. This study ficult decisions about where to allocate funds illustrates how different management actions and efforts. In many cases, we will have to will have different impacts, depending on the choose which populations, and perhaps even way climate changes in the future. Nonetheless, species, to let go extinct. As the ecological ef- it also demonstrates the possibility of identifying fects of climate change grow in magnitude in management options that yield benefits across response to more rapid changes in climate and all scenarios—in other words, “robust” options. disturbance regimes, triage will likely become Lawler: Climate Change Adaptation Strategies 91

Figure 3. Triage classification schemes for medical emergencies and for managing ecosystems and species in a changing climate. an integral part of higher-level planning and able resources. Others will need to be managed management. immediately and constantly, but such manage- Triage—from the French verb trier, to sort— ment will be feasible. Still others will need to be is a method of prioritization developed for treat- managed but might be able to wait a few years ing patients in emergency situations. Priority if they are closely monitored. Finally, the rest for treatment is based on the severity of the of the species and systems will either require injuries and the potential for survival. There no management or will require some manage- are generally four basic triage classifications: ment in the future, but won’t be lost if action deceased or expectant, critical, severe, and mi- isn’t taken soon. These systems and species can nor. Patients in each of these categories receive be monitored if resources are available. a different treatment (Fig. 3). Applying triage Medical triage approaches are often mod- to conservation and management decisions is ified to address specific situations. For exam- not a popular topic, and as such, has received ple, in a widespread emergency, if some of the relatively little attention. There are both ethi- injured include members of the medical pro- cal and ecological reasons that triage is a bitter fession, those doctors or nurses may receive a pill for both conservationists and conservation higher priority, even if they have relatively mi- biologists to swallow (Kareiva and Levin 2003). nor injuries, due to their value to the emergency The loss of a population or a species may have response effort. A triage system for addressing massive implications for the functioning of an climate change could likewise take the relative ecosystem in some cases or very little effect on importance of species or ecosystems into ac- a system in others. count (Fig. 4). Rare species or systems, or highly A simple triage system for addressing climate valued species might be deemed of higher pri- change would provide a classification based on ority even if the severity of climate impact severity that paralleled the medical classifica- was projected to be relatively low. Species with tion (Fig. 3). Some systems, species, or sites will high interaction strengths, particularly ecosys- likely undergo such substantial changes that tem engineers and keystone species, might, for they will be beyond managing with the avail- example, receive higher priorities than other 92 Annals of the New York Academy of Sciences

Figure 4. Triage classification for managing natural resources in a changing climate that accounts for both the severity of the climate threat and the value of the resource. The classifi- cation presented here is just one of a many potential manifestations of such a classification. species. For example, beaver have the ability to play a key role in prioritization and triage in to alter hydrological systems in ways that may the face of climate change. buffer streams from the effects of reduced snow- pack and earlier spring runoff events. Thus, even if beaver are not likely to be highly affected Future Research Directions by climate change, it may be wise to ensure the resilience of beaver populations in some systems The vast majority of recommendations for for the benefit of the system as a whole. Systems future research involve better understanding and species that provide critical ecosystem ser- how species and systems will respond to climate vices (again, beaver are a good example for change (Peters and Darling 1985; Kappelle et al. services such as flood control and trout) might 1999; Noss 2001; Schlesinger et al. 2001; Hulme also be given higher priorities. There are clearly 2005; Root and Schneider 2006). Although other ways to evaluate species and ecosystems such an understanding is clearly important, di- that go beyond ecological impact. For exam- recting the bulk of our research efforts to this ple, the loss of a species may have little impact goal will likely produce far too little, much too ecologically, but a large impact socially or eco- late. Because responses to climate change will nomically (Ruckelshaus et al. 2003). Here again be species and system specific, we will likely the concept of ecosystem services may be able discover few new generalities that will allow us Lawler: Climate Change Adaptation Strategies 93 to develop widely applicable adaptation strate- needs involves gaining a better understand- gies. Arguably, many of these generalities are ing of how to implement adaptive manage- already known (e.g., species will move in re- ment. What is the best way to explore multiple sponse to climate change, changes in phenol- climate-change scenarios in an adaptive man- ogy will decouple ecological systems). Instead agement setting? What will need to be moni- of solely attempting to document system- and tored? How often will monitoring need to be species-specific responses to climate change, to done? There has long been a call for increas- successfully address the challenge of how to re- ing the amount of adaptive management that is spond to these changes, research efforts will actually implemented. Implementing adaptive need to have a much more applied focus. management in a changing climate will both Because understanding how each species allow us to learn more about the ecological ef- and system will respond to climate change is fects of climate change and to provide flexible not feasible, perhaps the most critical task for management approaches. researchers is to determine which, if any, of Assigning priorities will require an under- the general concepts and basic prescriptions for standing of which systems and species will be adaptation will work and where and how they most vulnerable to climate change. Assessments will work best. With respect to conservation of climate-change vulnerability and potential planning, we should be asking whether aligning impacts can provide that knowledge (Desanker reserves along latitudinal or elevational gradi- and Justice 2001; Kareiva et al. 2008). Although ents, connecting reserves with stepping stones, there are many potential approaches to de- and expanding the bioclimatic footprint of re- veloping a vulnerability assessment for climate serves will provide more protection than merely change, any assessment would need to include increasing the number or size of reserves. With at least two basic elements: (1) a measure of in- respect to removing other threats and envi- herent sensitivity,and (2) projections of how and ronmental stresses, we will need to determine where climate will change. Additionally, a vul- which existing threats and stresses will have nerability assessment might include estimates the strongest synergistic interactions with ris- of the adaptive capacity of a system, species, ing temperatures and changing precipitation management agency, or society in general regimes. (McClanahan et al. 2008). Measures of inher- Many of the general adaptation strategies ent sensitivity can be taken from the literature, focus on managing systems and landscapes to although, for many species and systems there is either lessen the impacts of climate change or still little knowledge about their sensitivities to to allow species to move in response to climate changes in temperature or precipitation. The change. Fewer strategies are aimed at increas- vulnerability of individual species could be de- ing the potential for evolution in the face of termined by factors such as physiology, specific climate change. For many long-lived species, habitat requirements, interspecific dependen- evolution will not be an option. Others, how- cies, dispersal ability, and population dynam- ever, may be able to evolve in response to ics and location. An assessment of ecosystem warmer temperatures, wetter or drier condi- vulnerability would include factors such as hy- tions, or changing habitats. Which populations drologic or fire sensitivities, component–species and which species will have more evolutionary sensitivities, and vulnerability to sea-level potential? How can we promote evolutionary rise. potential through restoration, management, Determining where the largest climatic and conservation planning? changes are likely to occur will require fine- Given the critical role that adaptive man- scale predictions of future climate. There are agement is likely to play in addressing climate several methods that take local climatic condi- change, one of the most important research tions and topographic variation into account to 94 Annals of the New York Academy of Sciences derive finer resolution climate data from coarse Conclusions resolution general circulation model (GCM) projections (Wilby et al. 1998). Because of the Managers already have many of the tools variability across GCM projections, it will be necessary to address climate change. The vast necessary to assess potential future climatic majority of these tools are those recommended changes based on a range of different climate- for protecting biodiversity and managing nat- change projections. In addition to assessing cli- ural resources in general. What is needed in matic change, it will often be necessary to as- the face of climate change is a new perspec- sess where potential climate-driven changes in tive. Each of these tools—enhancing connec- disturbance regimes, the structure of vegeta- tivity, restoration, translocations—will need to tion, and species composition will be the great- be applied in light of the fact that disturbance est. Hydrological models can be used to gen- regimes and ecosystems will change and species erate predicted changes in hydrology (de Wit and pathogens will move. This new perspec- and Stankiewicz 2006), and fire models can be tive will require expanding the spatial and used to project changes in fire frequency, size, temporal scale of management and planning. and severity (McKenzie et al. 2004). DGVMs It will require restoring ecosystem function- can be used to generate predicted shifts in ing and managing for ecosystem services in- basic vegetation types corresponding to gen- stead of species composition. It will include eral habitat types (Bachelet et al. 2001; Cramer active adaptive management using scenario- et al. 2001) and climate-envelope models can be based approaches. And, it will involve priori- used to assess how particular, sensitive species tization, and as climatic impacts become more will likely respond to climate change or where acute, triage to determine which species and changes in flora or fauna might be the great- which ecosystem processes we will save. est (Thuiller et al. 2005; Lawler et al. in press- a). Ideally, these species distribution models should take species dispersal abilities and land- Acknowledgments scape patterns into account to determine how species will move and where greenways and I thank Molly Cross, Peter Kareiva, and Lara habitat corridors might be placed to enhance Hansen for stimulating discussions that helped movement. shape the paper. Molly Cross, Evan Girvetz, Finally, understanding how climate change Peter Kareiva, and Bill Schlesinger provided will affect ecosystem services will be critical helpful suggestions for improving earlier drafts. for setting priorities, conducting triage, and de- signing restoration projects. As discussed ear- Conflicts of Interest lier in this chapter, assigning limited resources to a large number of potential management The author declares no conflicts of interest. projects in an efficient way will require prioritiz- ing those projects based on both the severity of potential climate impacts and the values of the References systems, species, or populations. The concept of ecosystem services is one such valuation ap- Araujo,´ M.B., M. Cabeza, et al. 2004. Would climate proach. Moving from species-based restoration change drive species out of reserves? An assessment to a concentration on restoring ecosystem func- of existing reserve-selection methods. 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